issn 1239-6095 (print) issn 1797-2469 (online) helsinki 28 June 2013
Editor in charge of this article: Eeva-Stiina Tuittila
atmospheric impact of abandoned boreal organic agricultural soils depends on hydrological conditions
marja maljanen
1)*, Jyrki hytönen
2), Päivi mäkiranta
3), Jukka laine
4), Kari minkkinen
3)and Pertti J. martikainen
1)1) University of Eastern Finland, Department of Environmental Science, P.O. Box 1627, FI-70211 Kuopio, Finland (*corresponding author’s e-mail: marja.maljanen@uef.fi)
2) Finnish Forest Research Institute, Silmäjärventie 2, FI-69100 Kannus, Finland
3) Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland
4) Finnish Forest Research Institute, Kaironiementie 54, FI-39700 Parkano, Finland Received 8 July 2011, final version received 16 Mar. 2012, accepted 13 Mar. 2012
maljanen, m., hytönen, J., mäkiranta, P., laine, J., minkkinen, K. & martikainen, P. J. 2013: atmos- pheric impact of abandoned boreal organic agricultural soils depends on hydrological conditions.
Boreal Env. Res. 18: 250–268.
Drained agricultural peat soils are significant sources of carbon dioxide (CO2) but also small sinks for methane (CH4). Leaving these soils without any cultivation practice could be an option to mitigate GHG emissions. To test this hypothesis, we measured, over a three year period, net CO2 exchange and fluxes of CH4 for five agricultural peat soils that had been abandoned for 20–30 years. Annually, the sites were either small net sinks or sources of CO2 and CH4 (–7.8 to 530 g CO2-C m–2 and –0.41 to 1.8 g CH4 m–2). Including N2O emissions from our previous study, the net (CH4 + CO2 + N2O) emissions as CO2 equiva- lents were lower than in cultivated peat soils and were lowest in the wet year. Therefore, high GHG emissions from these soils could be avoided if the water table is maintained close to the soil surface when photosynthesis is favoured over respiration.
Introduction
Pristine, water-saturated peat soils store large amounts of carbon because the rate of biomass production in photosynthesis is greater than the rate of decomposition (Gorham 1991, Alm et al.
1997, Turunen 2008). The carbon accumulation rate in undrained boreal and subarctic peatlands is, according to Turunen and Moore (2003), 13–20 g m–2 yr–1. Pristine peatlands have negligi- ble emissions of nitrous oxide (N2O), (Martikai- nen et al. 1993), but they are sources of methane (CH4) (Lai 2009, Saarnio et al. 2009). Drain- age, fertilization, ploughing, irrigation, harvest- ing, liming and compaction of soil by machines
modify the soil properties of managed peatlands and thereby change their carbon and nitrogen cycles (e.g. Martikainen et al. 1993, Regina et al.
1998, Lohila et al. 2004, Ball et al. 2008). Man- aged boreal peat soils are generally substantial net sources of CO2 (Kasimir-Klemedtsson et al.
1997, Mosier et al. 1998, Maljanen et al. 2001, 2004, Lohila et al. 2004) and N2O (Kasimir- Klemedtsson et al. 1997, 2009, Maljanen et al.
2003a, Regina et al. 2004, Rochette et al. 2010) but low sources, or even sinks, for atmospheric CH4 (Kasimir-Klemedtsson et al. 1997, 2009, Maljanen et al. 2003b, Regina et al. 2007).
The use of organic soils in agriculture is among the most problematic land-use options
when considering atmospheric impact. For example, in Finland 8% of the total greenhouse gas (GHG) emissions originate from agricul- ture. Agricultural soils cover about 6% of the land area in Finland. Organic agricultural soils cover only 13% of the agricultural soils (Myllys and Sinkkonen 2004) and 0.6% of the total peatland area in Finland (Turunen 2008), but they are mainly responsible of the GHG emis- sions originating from agriculture (Statistics Fin- land 2010). In Sweden, 6%–8% of total GHG emissions originate from cultivated peatlands which represents 8.6% of the total arable land area (Berglund and Berglund 2010). Therefore, an important question is how to reduce the GHG emissions from these managed organic soils covering only a relatively small area. One option would be to cease all agricultural activi- ties like ploughing, fertilization, harvesting on these problematic soils, but how this would change the GHG emissions from these soils is poorly understood. Cropland ecosystems lose large amounts of nutrients with the harvested crop each year, which must be replaced by fer- tilization. Peat soil also needs to be limed to keep the soil pH high enough for crops (Myllys 1996). Secondary vegetation succession start- ing after abandonment could incorporate more carbon into the ecosystem. Grasses and herbs dominate in field vegetation for decades during secondary succession (Törmälä 1982), and open ditches are the first habitat for pioneer tree spe- cies (birch, willows) (Hytönen 1999). In aban- doned systems, there could thus be an increase in the energy flow to decomposers (Törmälä 1982).
On the other hand, a decrease in soil pH without regular liming could reduce the decomposition rate. Decomposition and CO2 emissions could also decrease as a result of lower soil aeration after ploughing has ended, and after increase in water-table level if the ditch network is not maintained any more after abandoning the site (Yavitt et al. 1997).
We hypothesize that when former organic agricultural soils are maintained without any management for several decades, they lose less CO2 than cultivated peat soils as a result of a reduction in the soil respiration rate and increased photosynthesis after plant succes- sion. After gradual deterioration of the drainage
ditches, a rise in the water-table level would decrease decomposition of soil organic matter and favour carbon accumulation in the ecosys- tem. However, there is a risk that CH4 emissions would increase with the increasing water-table level (e.g. Tuittila et al. 2000). To study these hypotheses, we measured annual CO2 balances and CH4 fluxes from five abandoned agricultural soils for three years which, along with previous data on their N2O fluxes (Maljanen et al. 2012), completes the picture of changes in GHG emis- sions.
Material and methods
Study sites
CO2 exchange and CH4 fluxes were measured from June 2002 to June 2005 for five aban- doned organic agricultural fields (AB1–AB5, Table 1). The fields were located within 700 m of each other in Kannus, western Finland (63°54´N, 23°56´E). The length of the growing season in Kannus is about 180 days and the soil is usually covered with snow from November to mid-April.
The mean annual temperature is 2.4 °C and the annual precipitation is 561 mm. The sites, originally pristine peatlands, had been drained in the 1950s and used for cultivation of perennial grasses and cereals (crop rotation) for decades before they were abandoned 20–30 years prior to the current study. The crop rotation (perennial grasses/cereals) is a normal practise in Finland.
During the agricultural phase, some mineral soil had been mixed with the peat, and normal NPK fertilization (40–100 kg ha–1 N, 17–35 kg ha–1 P and 33–80 kg ha–1 K according to Lampinen 1978 and Rainiko 1978) was used in order to improve the soil properties for cultivation. No fertilization or ploughing activities have been carried out since the agricultural use ended, and the sites were naturally vegetated. The dominant plant species (mean coverage more than 10%) were Juncus filiformis and Deschampsia cespi- tosa at sites AB1 and AB2; Elymus repens and Poa pratensis at site AB3; Deschampsia cespi- tosa, Epilobium angustifolium and Rubus arcti- cus at site AB4 and Epilobium angustifolium, Deschampsia cespitosa and Agrostis capillaris
at site AB5. There were no trees growing in the middle of the fields, but some birch and willow shrubs were growing by the ditches.
Soil chemical and physical characteristics and weather data
At each site, five soil samples (239 cm3) were taken from the depths of 0–10 and 10–20 cm.
Soil pH(H2O) was measured from dried soil sam- ples using a 1:2.5 v:v soil:solution suspension.
The bulk density of the soil was calculated as the ratio of dry mass (dried at 105 °C) to volume of the sample. The organic matter content was determined as loss-on-ignition (at 550 °C for 8 h). The content of total soil C was determined with a LECO CHN-1000 or LECO CHN-2000.
The total N of the soil samples was determined by the Kjeldahl method, and the total concentra- tions of P and K using HCl extraction of ignition residues. Decomposition status of the peat was determined according to the von Post humifica- tion scale (von Post 1922).
Temperatures of the unfrozen soil were meas- ured manually (Fluke 51 II Digital Thermom- eter) at the depths of 2, 5, 10 and 20 cm during gas sampling. At site AB3 (located in the middle of the sites), soil temperature (depths 5 and 20 cm) was monitored continuously using Camp- bell 107 soil temperature sensors and a data logger. Air temperature and precipitation data were collected by the Finnish Meteorological Institute at Toholampi weather station near the study sites (Fig. 1 and Table 2). The water-table level (WT) was measured in groundwater wells,
and the depth of soil frost was measured at each site using frost-depth gauges filled with methyl- ene blue–H2O solution. The distance of WT from the soil surface is indicated by a negative value.
The cellulose decomposition test was carried out during two summers and one winter from 6 June to 11 September 2003 (98 d), from 15 Octo- ber 2003 to 7 May 2004 (205 d), and from 7 May to 15 September 2004 (131 d). Pieces of birch- wood cellulose (5 ¥ 10 cm) were dried at 105 °C, stabilized for two hours at room temperature and weighed. Three pieces were inserted into a plas- tic net (mesh size 1 mm), and three replicate nets were buried in the peat at each site at the depths of 0–5, 5–10 and 10–15 cm. After the in situ incubation period, the ingrown roots and mosses were cleaned off, and the pieces were dried and weighed and the cellulose decomposition rate was calculated from the weight loss.
Plant cover characteristics
The total coverage of the understorey plants and the composition of the dominant plant species were recorded in early August 2002, 2003 and 2004. Above-ground plant biomass was collected at the end of July 2003 and 2004 using a 20 ¥ 20 cm frame with six replicate cuttings. Samples were dried at 105 °C and the amount of biomass (g m–2) was calculated. The growth dynamics of the predominant plant species (Deschampsia cespitosa, Elymus repens, Agrostis capillaris and Juncus filiformis) were measured using marked plants, whose size and number of leaves were determined. Leaf area (LA) was calculated by
Table 1. site characteristics measured in 2002 (soil sampling depth 0–20 cm). PD = peat depth, BD = bulk density, om = organic matter content, c = carbon, n = nitrogen, cD = cellulose decomposition, Ptot = total phosphorous, Ktot
= total potassium and ph = measured from 1:2.5 v:v soil:h2o suspension, H = degree of decomposition (von Post 1922).
site PD BD om c:n n cDa) Ptot Ktot phwater H
(cm) (g cm–3) (%) (%) (% d–1) (mg g–1) (mg g–1)
aB1 30 0.38 52 18.8 1.2 0.30 1.5 1.6 4.9 8–9
aB2 20–30 0.40 43 19.0 0.9 0.32 0.1 1.1 5.0 8–9
aB3 30 0.38 44 18.3 1.0 0.25 0.2 1.7 5.9 8–9
aB4 30 0.30 61 16.3 1.7 0.33 0.2 0.7 4.5 9
aB5 80 0.42 41 19.2 1.1 0.47 0.1 2.5 4.3 10
a) mean value from three periods in 2003–2004.
Temperature (°C)
–40 –30 –20 –10 0 10 20 30
Tair T5 T20
Precipitation (mm)
0 10 20 30 40
50 Precipitation
Day number 120
PAR (µE s–1)
0 200 400 600
PAR a
c b
180 240 300 360 420 480 540 600 660 720 780 840 900 960 1020 1080 1140 1200
Fig. 1. (a) Daily mean soil (at 5 and 20 cm depth) and air temperatures at the study site aB3, (b) daily precipitation, and (c) mean daily photosynthetically-active radiation (Par) at the toholampi weather station.
multiplying the number of leaves inside the frames by their length and width (Wilson et al.
2007). The coverage of all other plants measured in the late summer was used to estimate the total
leaf area inside the frames. LA was measured six times during the growing season in 2002, and 12 times in 2003 and 2004, and the gaps between the measurements were interpolated.
CO2 exchange and CH4 fluxes during the growing season
Weekly CO2 exchange measurements were carried out from July 2002 until October 2004 during the growing seasons using a chamber method (Alm et al. 1997). The instantaneous net CO2 exchange was measured with a transparent chamber (60 ¥ 60 cm, height 30 cm) with a thermostat, which was placed over an aluminium frame (58 ¥ 58 cm) pre-installed in the soil. There were four rep- licates for each site. To ensure gas tightness, there was a groove in the upper edge of the frame which was filled with water. A portable infrared gas ana- lyzer (EGM 3 or EGM 4, PP Systems, USA) was used to measure the change in the CO2 concentra- tion in the chamber during an incubation period of three minutes under stable irradiation conditions.
CO2 concentration readings were recorded at 15-s intervals. After a measurement with full irradia- tion, the chamber was shaded with one or two mesh fabrics, and the measurement was repeated under this reduced irradiation. The light intensity (PAR) and air temperature (Ta) inside the chamber were recorded during the incubation period. The total respiration (Reco) of the plant–soil system was measured as described above in the dark by covering the chamber with an opaque lid. In addi- tion to the four vegetated plots, one frame at each site was kept free of above-ground vegetation in order to measure soil respiration (Rsoil). The CO2 flux was calculated from the linear part of the CO2 concentration slope in the light and in the dark (Alm et al. 1997).
CO2 uptake from the atmosphere to the eco- system is indicated here with a negative, and release of CO2 to the atmosphere with a positive
sign. An estimate of GPP (gross primary produc- tion) was calculated as the sum of CO2 fluxes measured under light and dark conditions (Alm et al. 1997). The measurement in the dark was used as an estimate of the total respiration (Reco), and net ecosystem exchange (NEE) was then calculated as:
NEE = GPP – Reco (1)
For calculation of the diurnal NEE values, statistical response functions were constructed separately for each site for the growing seasons of 2002, 2003 and 2004 in order to reconstruct the hourly values for GPP and Reco on the basis of climatic data (Table 3). The response functions for GPP and Reco or Rsoil were as follows:
GPP = Q ¥ I ¥ LA/(k + I) (2) Reco or Rsoil = exp(b0 + b1T5 + b2WT) (3) The dependence of GPP on irradiation, I, (µmol m–2 s–1) takes the form of a rectangular hyperbola function (e.g. Frolking et al. 1998), where Q is the asymptotic maximum, and k is the half saturation constant. The additional parame- ter LA in Eq. 2 is the leaf area (cm2 cm–2). In Eq.
3, T5 is the soil temperature at the depth of 5 cm, and WT is the water-table depth (cm) assuming negative values when the water table is below soil surface. The diurnal estimates of GPP and Reco were reconstructed using hourly time-series data for T5, LA, WT and I from the beginning of May until the end of October each year.
CH4 fluxes were measured throughout the three years, every second or third week (Fig. 2).
During the snow-free periods, fluxes of CH4
Table 2. the mean seasonal temperatures and precipitation (from 1 may to 31 october and from 1 november to 30 april) in Kannus, and their comparison (%) to the long term average (1971–2000) (Drebs et al. 2002).
may–october november–april
mean temp. Precipit. mean temp. Precipit.1)
(°c) (%) (mm) (%) (°c) (%) (mm) (%)
2002–2003 10.3 101 283 81 –7.7 145 190 90
2003–2004 9.6 94 479 137 –4.3 81 241 115
2004–2005 10.5 103 555 155 –3.5 66 247 118
1) mostly as snow.
Table 3. Parameters for the response functions to calculate gross primary photosynthesis (GPP), ecosystem respi- ration (Reco) and soil respiration (Rsoil) for the growing seasons of 2002, 2003 and 2004. Units: mg m–2 h–1 for GPP, Reco, Rsoil; µs m–2 for I; cm2 cm–2 for la; °c for T, and cm for Wt (negative values when below soil surface).
site aB1 aB2 aB3 aB4 aB5
GPP = [QI/(k + I )] ¥ la 2002
Q 361 ± 43.2 529 ± 62.3 191 ± 19.9 333 ± 40.5 251 ± 59.4
k 269 ± 91.4 252 ± 90.1 188 ± 66.1 272 ± 89.1 340 ± 214
r 2 0.74 0.65 0.62 0.69 0.42
2003
Q 154 ± 12.7 422 ± 44.6 403 ± 55.5 493 ± 58.7 565 ± 45.8
k 261 ± 69.9 369 ± 101 463 ± 148 263 ± 91.6 163 ± 49.7
r 2 0.67 0.51 0.34 0.23 0.56
2004
Q 307 ± 29 384 ± 28.8 440 ± 41.1 243 ± 25.5 388 ± 34.3
k 400 ± 88.8 198 ± 51.7 285 ± 82.2 337 ± 89.9 461 ± 96.6
r 2 0.48 0.47 0.56 0.18 0.67
lnReco = b0 + b1¥ T5 + b2¥ Wt 2002
b0 5.30 ± 0.07 5.56 ± 0.12 5.99 ± 0.06 5.43 ± 0.16 5.82 ± 0.13
b1 0.11 ± 0.01 0.06 ± 0.01 0.07 ± 0.01 0.05 ± 0.01 0.04 ± 0.01
b2 0.002 ± 0.001 –0.01 ± 0.002 – –0.01 ± 0.003 –0.004 ± 0.002
r 2 0.88 0.80 0.79 0.73 0.48
2003
b0 5.67 ± 0.11 6.13 ± 0.10 5.89 ± 0.12 6.54 ± 0.07 5.78 ± 0.12
b1 0.05 ± 0.01 0.04 ± 0.01 0.08 ± 0.01 – –0.07 ± 0.01
b2 –0.01 ± 0.002 –0.01 ± 0.001 –0.01 ± 0.002 –0.01 ± 0.001 –0.01 ± 0.002
r 2 0.80 0.74 0.74 0.52 0.62
2004
b0 5.26 ± 0.16 5.44 ± 0.12 5.61 ± 0.12 5.77 ± 0.11 5.24 ± 0.15
b1 0.12 ± 0.01 0.12 ± 0.01 0.13 ± 0.01 0.10 ± 0.01 0.12 ± 0.01
b2 –0.02 ± 0.004 –0.01 ± 0.002 –0.01 ± 0.002 –0.01 ± 0.002 –0.01 ± 0.002
r 2 0.51 0.69 0.72 0.68 0.72
lnRsoil= b0 + b1¥ T5 + b2¥ Wt
b20020 4.96 ± 0.18 5.56 ± 0.78 4.24 ± 1.45 4.81 ± 0.59 4.59 ± 0.69 b1 0.06 ± 0.02 –0.004 ± 0.04 0.03 ± 0.05 0.04 ± 0.03 0.01 ± 0.04 b2 –0.01 ± 0.003 –0.01 ± 0.01 –0.01 ± 0.02 –0.01 ± 0.01 –0.01 ± 0.01
r 2 0.93 0.37 0.46 0.78 0.45
b20030 4.30 ± 0.56 5.14 ± 0.40 4.79 ± 0.32 4.74 ± 0.41 3.98 ± 0.30
b1 0.06 ± 0.04 0.05 ± 0.01 0.07 ± 0.02 0.03 ± 0.03 0.09 ± 0.02
b2 –0.03 ± 0.01 –0.01 ± 0.001 –0.01 ± 0.004 –0.02 ± 0.01 –0.01 ± 0.004
r 2 0.80 0.61 0.79 0.73 0.83
b20040 3.23 ± 0.47 3.39 ± 0.25 3.14 ± 0.54 3.81 ± 0.11 3.77 ± 0.11
b1 0.12 ± 0.03 0.12 ± 0.02 0.10 ± 0.04 0.08 ± 0.03 0.18 ± 0.02
b2 –0.04 ± 0.02 –0.03 ± 0.01 –0.04 ± 0.01 –0.04 ± 0.002 –0.01 ± 0.01
r 2 0.61 0.89 0.70 0.70 0.90
were measured with a static chamber method using aluminium frames as described above, and an opaque aluminium chamber (60 ¥ 60 cm, height 30 cm) equipped with a fan. This method
was used also for CO2 (Reco) outside growing season during snow free periods. Gas samples (40 ml) were drawn from the headspace of the chambers with 60 ml polypropylene syringes
0 1 2 3
–120 –100 –80 –60 –40 –20 0
–0.1 0 0.1 0.2
–120 –100 –80 –60 –40 –20 0
CH4 flux (mg CH4-C m–2 h–1) –0.1
0 0.1 0.2
WT (cm)
–120 –100 –80 –60 –40 –20 0
–0.1 0 0.1 0.2
–120 –100 –80 –60 –40 –20 0
Day number
120 180 240 300 360 420 480 540 600 660 720 780 840 900 960 1020 1080 1140 1200 –0.2
–0.1 0 0.1 0.2
–120 –100 –80 –60 –40 –20 0 AB1
AB5 AB4 AB3 AB2
CH4 flux WT
Fig. 2. methane (circles) dynamics (mg ch4-c m–2 h–1) and water table depth (Wt, distance from the soil surface, dots) on abandoned organic agricultural soils from July 2002 to June 2005. negative and positive flux values indi- cate uptake and emissions of ch4 by the ecosystem, respectively.
(Becton Dickinson) equipped with three-way stopcocks 5, 10, 15 and 25 minutes after the chambers were installed on the frames. The sam- ples were analyzed with a gas chromatograph (Shimadzu GC-14B or Hewlett-Packard 5890) equipped with a flame ionization (FI) detector for CH4, and thermal conductivity (TC) detec- tor for CO2 (Maljanen et al. 2001). Standards of 1.98 µl l–1 CH4 and 396 µl l–1 CO2 were used for hourly calibrations. The gas flux rates were cal- culated from the linear change in the gas concen- trations in the headspace of the chamber.
CH4 and CO2 fluxes during winter with snow
During winter, when the snowpack depth was more than 15 cm, the CH4 and CO2 fluxes were determined using a gas gradient technique (Som- merfeld et al. 1993). Gas samples for the concen- tration analyses were drawn from the snow pack into the syringes using a stainless steel probe (diam. 3 mm). Samples were taken inside the collars and from several depths from 2 cm above the soil surface to 5 cm below the snow surface to verify linearity. Gas concentrations were ana- lysed within 24 hours of sampling with a gas chromatograph as described above. Annual CH4 and CO2 flux rates during the winter period were calculated for each site separately using time- weighted average values for measured emissions.
Additional gas flux data and GWP calculations
Published N2O flux data from the same sites and same period (Maljanen et al. 2012) were used to calculate the total atmospheric impact of CO2, CH4 and N2O fluxes as CO2 equivalents using the GWP approach (100 year time horizon) accord- ing to Solomon et al. (2007). The annual emis- sions of CH4 were multiplied by 25 and those of N2O by 298 to convert these emissions to CO2 equivalents. Similar calculations were made for published data from grass leys with low man- agement intensity (Maljanen et al. 2001, 2003a, 2003b, 2004, Lohila et al. 2004, Regina et al.
2004, 2007) and cultivated barley fields with fre-
quent management (Maljanen et al. 2001, 2003a, 2003b, 2004) on organic soils in Finland. The grass leys were ploughed and sown every 3–4 years and the barley fields annually; both were also fertilized (NPK) annually.
Statistical methods
Parameters for the photosynthesis functions were calculated using a non-linear regression, and coefficients for the respiration functions using a stepwise linear regression. CH4 flux rates were not normally distributed and there- fore correlations between the CH4 flux rates and environmental parameters were calculated using Spearman’s rank correlation (rS). The differences in cellulose decomposition rates were tested with a paired t-test. All tests were performed using SPSS ver. 17 (SPSS Inc.)
Results
Soil characteristics
At the depth of 0–20 cm, soil pH(H2O) varied from 4.3 to 5.9, and the soil C:N ratio from 16.3 to 19.2 (mean = 18.3). All the soils were rather well decomposed, in von Post scale (H) from 8 to 10 (Table 1). The organic matter content in the same peat layer varied from 41% to 61%, and the peat bulk density between 0.30 and 0.42 g cm–3 (Table 1). The mean cellulose decom- position rate (average from all depths) varied between the sites from 0.18% d–1 to 0.67% d–1 during the summer periods (burial times 98 to 131 days) and from 0.02% to 0.33% during the winter (burial time 205 days). Cellulose mean decomposition values in 2003 and in 2004 were 0.55% d–1 and 0.34% d–1, respectively, but they did not differ statistically.
Weather conditions
The 2002 growing season was warm and dry.
The average temperature during the summer months (from June to August) was 16.0 °C and the maximum temperature, measured in July,
was 28 °C. The following winter was cold;
the lowest air temperature recorded at the end of December 2002 was –40 °C (Fig. 1a). The 2003 growing season was cooler and wetter; the average temperature from June to August was 13.6 °C. The following winter was milder; tem- peratures dropped to below –20 °C only a few times. The third growing season (2004) was wet (Fig. 1b) and slightly warmer than that of 2003.
The mean temperature from June to August was 13.8 °C and the precipitation (304 mm) was 100 mm above the long-term average (for years 1971 to 2000, Drebs et al. 2002). The following winter was milder; air temperature remained above –20 °C. The mean temperatures and pre- cipitation levels during the growing seasons (from May to October) and winters (November to April) are listed in Table 2.
In 2002, soils started to freeze early in the late November but in the following years, soil frost developed later. The maximum soil-frost depth varied from 4 to 17 cm. Soils thawed between late April and early May. The maximum soil temperature, 19.6 °C (at a depth of 5 cm), during the three year period was measured in July 2003 and the minimum, –2.4 °C, in March 2004. During winter, soil temperature remained close to –0.5 °C at all sites (Fig. 1a).
The water-table (WT) level was lowest during the dry, first growing season, and high- est during the wet, third growing season. The WT varied from 128 cm below the soil surface to 5 cm above the soil surface, being on average 46 cm below the soil surface between May and October (Fig. 2).
Vegetation characteristics
Predominant plant species varied between the sites. The total coverage of plants in the third growing season was close to 90%, whereas in the first growing season it was clearly lower (Table 4). The maximum leaf area in the study plots was also lowest in 2002 and highest in 2004. Plant biomass was not measured in the first growing season but it was greater in the wet, third growing season than in the previous year. The above-ground biomass varied from 27 to 269 g C m–2. The below-ground biomass was
not measured, but the total biomass for peren- nial grasses was estimated to be 2.5 times their above-ground biomass (Törmälä 1982, Pietola and Alakukku 2005), and it ranged from 38 to 376 g C m–2 (Table 4).
Ecosystem respiration (Reco) and soil respiration rates (Rsoil)
The maximum Reco values during the growing season were about 1000 mg CO2-C m–2 h–1, whereas during winter, the respiration rates were low, from 20 to 50 mg CO2-C m–2 h–1 (Fig. 3).
The Reco during the growing season correlated with soil temperature (at 5 cm depth) and WT level and, therefore, these variables were used in the response functions (Table 3) for calculating diurnal Reco. The modelled Reco rates were at their lowest during the dry growing season (184 days) of 2002 (from 831 to 1057 g CO2-C m–2), high- est in 2003 (from 937 to 1570 g CO2-C m–2) and slightly lower during the 2004 growing season (from 760 to 1460 g CO2-C m–2). Winter respi- ration rates (from 1 November to 30 April, 181 days) were from 92 to 221 g CO2-C m–2.
Soil respiration rates (Rsoil) were on average of 50%, 40% and 20% of Reco during the first, second and third growing seasons, respectively.
The annual modelled Rsoil rates were at their highest during the second year and at their lowest during the wet third year (Table 4).
CO2 exchange and CO2 balance
The gross primary production (GPP) was strongly dependent on photosynthetically active radiation (I ) during the growing season. Leaf area also correlated with GPP and was used in the response function as described above (Table 3). The modelled GPP rates during the growing seasons ranged from 642 to 1094 g CO2-C m–2 (Table 4 and Fig. 4). The modelled GPP values were highest during the wet growing season of 2004 and lowest during the dry and warm growing season of 2002. GPP increased with higher WT levels (r2 = 0.84, 0.97, 0.99, 0.95 and 0.99 at sites AB1–AB5, respectively), thus high soil moisture favoured photosynthesis
(Fig. 5 and Table 5). Calculated instantaneous GPP values (measured Reco + NEE) in full irra- diation conditions were significantly higher at high WT level than at low WT level (Table 5).
There was also a positive correlation between the precipitation sum and GPP. When precipita- tion during the growing season increased from 300 mm (dry year) to 550 mm (wet year), GPP increased on average by a factor of 1.3. The total (above + below ground) biomass measured out- side the gas sampling plots in 2003 and 2004 did not correlate well with the GPP or NEE.
The modelled net ecosystem CO2 exchange during the growing seasons varied from a net sink of –122 g CO2-C m–2 to a net emis- sion of 383 g CO2-C m–2. When the winter emissions were included, the annual net CO2
exchange ranged from –7.8 (sink) to 530 (emis- sion) g CO2-C m–2 yr–1 (Table 4). Based on the three-year mean (mean ± SD = 306 ± 161 g CO2-C m–2), the ecosystems were annual net sources of CO2.
CH4 flux rates
The sites were either weak sinks for or sources of CH4, with an average (± SD) rate for the five sites of –0.018 ± 0.516 g CH4-C m–2 yr–1. Most of the sites acted as small sinks for CH4 during the first and second years. During the third year, site AB1 emitted 1.8 g CH4-C m–2 and site AB3 emitted 0.03 g CH4-C m–2 as a result of high water-table levels (Fig. 2 and Table 6). The other
Table 4. summary of mean water table depth (Wt), annual soil respiration (Rsoil), ecosystem respiration (Reco), gross primary production (GPP) and net ecosystem co2 exchange (nee) during three years (June 2002–may 2003, June 2003–may 2004 and June 2004–may 2005). negative sign indicates co2 uptake by the ecosystem and positive sign its emission to the atmosphere. above-soil biomass (sBm) was measured in the middle of July and the total biomass (Bm) was estimated as c. total coverage is the total coverage of plants inside the frames for the gas exchange measurements.
aB1 aB2 aB3 aB4 aB5 mean
2002–2003
Wt (cm) –70 –93 –109 –79 –91 –88
Rsoil (g co2-c m–2) 558 588 371 567 428 502
Reco (g co2-c m–2) 954 954 1179 939 1009 1007
GPP (g co2-c m–2) –666 –651 –840 –642 –606 –681
nee (g co2-c m–2) 287 303 339 297 403 326
total coverage (%) 42 43 85 48 65 57
2003–2004
Wt (cm) –29 –39 –58 –40 –55 –44
Rsoil (g co2-c m–2) 535 551 629 575 455 549
Reco (g co2-c m–2) 1030 1362 1606 1290 1332 1324
GPP (g co2-c m–2) –719 –832 –1094 –919 –833 –879
nee (g co2-c m–2) 310 530 511 325 499 445
sBm (g c) 55 27 104 55 30 54
Bm1) (g c) 77 38 145 77 42 145
total coverage (%) 55 55 86 67 62 65
2004–2005
Wt (cm) –2 –14 –28 –18 –35 –19
Rsoil (g co2-c m–2) 109 186 398 231 291 243
Reco (g co2-c m–2) 877 1115 1655 1289 1175 1222
GPP (g co2-c m–2) –885 –1000 –1205 –971 –1028 –1018
nee (g co2-c m–2) –7.8 115 450 318 147 204
sBm (g c) 194 269 203 134 137 187
Bm1) (g c) 272 377 284 188 192 262
total coverage(%) 80 80 100 94 85 88
1) estimated biomass c calculated from measured shoot biomass and with shoot:root ratio of 2.5:1 (according to Pietola and alakukku 2005).
AB2
–1500 –1000 –500 0 500 1000
AB1
–1500 –1000 –500 0 500 1000
RECO GPP NEE RECO
AB3
RECO, GPP, NEE (mg CO2-C m–2 h–1) –1500 –1000 –500 0 500 1000
AB4
–1500 –1000 –500 0 500 1000
AB5
–1500 –1000 –500 0 500 1000
Day number
120 180 240 300 360 420 480 540 600 660 720 780 840 900 960 1020 1080 1140 1200
Fig. 3. instantaneous co2 fluxes measured in all subplots using chambers and an irG analyzer. Black dots are ecosystem respiration rates (Reco) measured with an ir analyzer during the growing season and circles are eco- system respiration rates measured with Gc outside the growing season. triangles are net ecosystem exchange rates (nee) measured with a transparent chamber in a full light, and squares are calculated values of the gross primary production (GPP = nee + Reco). negative sign indicates uptake by the ecosystem and positive emission to the atmosphere.
AB1
–10 0 10 20
GPP NEE
AB2
–10 0 10 20
AB3
RECO, GPP, NEE (g CO2–C m–2 d–1) –10 0 10 20
AB4
–10 0 10 20
AB5
–20 –10 0 10 20
RECO
Day number
120 180 240 300 360 420 480 540 600 660 720 780 840 900 960 1020 1080 1140 1200
Fig. 4. modelled values of the ecosystem respiration (Reco, dashed line), gross primary production (GPP, solid thin line) and net ecosystem uptake (nee, solid thick line) for the abandoned organic agricultural soils during the grow- ing seasons of 2002, 2003 and 2004. negative sign indicates uptake by the ecosystem and positive emission to the atmosphere.
WT (cm)
–120 –100 –80 –60 –40 –20 0 GPP (g CO2-C m–2 yr–1)
–1300 –1200 –1100 –1000 –900 –800 –700 –600 –500
AB1 AB2 AB3 AB4 AB5 Fig. 5. Gross primary production (GPP) on abandoned agricultural peat soils during three growing seasons plotted against water table level (Wt) during the grow- ing season.
Table 5. mean gross photosynthesis ± sD (mg co2 m–2 h–1) measured under conditions of full irradiation (I > 700 µs s–1 m–2) and at the mean water-table level (cm, distance from the soil surface as a negative value) during meas- urement days at each site during the growing seasons of 2002, 2003 and 2004. the values with a common letter do not differ significantly at p ≥ 0.05 (tukey’s test).
mean GPP F, p Wt F, p
2002 2003 2004 2002 2003 2004
aB1 1660 ± 350a 2050 ± 850ab 2410 ± 670b 5.32, 0.007 –93a –46b –10c 98.3, < 0.001 aB2 1750 ± 320a 2170 ± 910ab 2470 ± 790b 3.72, 0.028 –118a –47b –28c 79.1, < 0.001 aB3 2130 ± 470a 2780 ± 1290a 3230 ± 1440a 2.78, 0.670 –136a –61b –34c 112, < 0.001 aB4 1970 ± 400a 2450 ± 1030ab 2850 ± 860b 3.91, 0.024 –113a –52b –26c 30.8, < 0.001 aB5 1870 ± 1170a 2250 ± 1120ab 2780 ± 1460b 4.32, 0.017 –128a –59b –40c 78.1, < 0.001
sites remained net sinks for CH4 during these years. The highest annual CH4 uptake rate (0.41 g CH4-C m–2) was measured at site AB5 in 2003 (Table 6).
Methane fluxes correlated with the WT level (Fig. 2) and soil temperature (at the depth of 5 cm) at all sites. CH4 uptake decreased with higher WT level (rS = –0.188, p = 0.030; rS = –0.444, p <
0.001; rS = –0.241, p = 0.004; rS = –0.292, p <
0.001; rS =–0.276, p = 0.002; at sites AB1 to AB5, respectively). CH4 uptake decreased with decreas- ing soil temperature at the depth of 5 cm (rS = 0.130, p = 0.046; rS = 0.332, p < 0.001; rS = 0.270, p < 0.001; rS = 0.268, p < 0.001; rS = 0.559, p <
0.001; at sites AB1 to AB5, respectively).
Table 6. annual net exchange of ch4, n2o and co2 (g m–2 yr–1) on abandoned organic agricultural soils during three years (June 2002–may 2003, June 2003–may 2004 and June 2004–may 2005). the GWPtot is the total warming effect (sum of ch4, co2 and n2o) as co2 equivalents. ch4 flux is multiplied by 24 and n2o flux by 298 to express fluxes as co2 equivalents (g co2 eq. m–2 yr–1) for 100-year time horizon (solomon et al. 2007). negative sign indi- cates uptake by the ecosystem and positive sign emission to the atmosphere.
site aB1 aB2 aB3 aB4 aB5 mean
2002–2003
ch4 –0.18 –0.23 –0.17 –0.20 –0.38 –0.23
co2 1050 1110 367 1090 1480 1020
n2o 0.54 0.99 0.43 0.73 2.45 1.03
GWPtot 1210 1400 490 1300 2200 1320
2003–2004
ch4 –0.12 –0.26 –0.12 –0.27 –0.55 –0.26
co2 1140 1940 1880 1190 1830 1960
n2o 0.28 0.51 0.69 0.85 3.17 1.10
GWPtot 1220 2090 2080 1440 2760 1920
2004–2005
ch4 2.41 –0.01 0.04 –0.08 –0.22 0.43
co2 –29 420 1650 1170 538 750
n2o 0.15 0.42 0.32 0.33 1.08 0.46
GWPtot 74 545 1750 1270 855 897
Total GWP of abandoned boreal organic agricultural soils
The global warming potential (GWP) of the sites, including C gases (CO2 + CH4), was calcu- lated as CO2 equivalents (Solomon et al. 2007).
These gases together had a warming effect of 28 to 1940 g CO2 eq. m–2 yr–1, with a mean of 1121 g CO2 eq. m–2 yr–1 (time horizon of 100 years).
Including N2O emissions from Maljanen et al.
(2012) resulted in a total mean (± SD) GWP of 1380 (± 677) g CO2 eq. m–2 yr–1 for these sites (Table 6).
Among the sites, AB1, for which the WT was close to the soil surface, had the lowest total GWP. The CO2 balance mainly determined the total GWP, whereas CH4 had only a minor effect (on average less than 0.1%, max. 3.2% of total GWP). The N2O emissions (Maljanen et al.
2012) increased the GWP on average by 21%.
Discussion
Weather and vegetation characteristics The three studied years differed greatly from each other with regard to precipitation and water table depth. The first growing season, 2002, was warm and dry; the second, 2003, cooler and wetter; and the third, 2004, very wet. Precipita- tion during the growing season of 2004 was almost twice the long-term average. Temperature and precipitation (mostly as snow) increased from the winter of 2002 to 2004. Vegetation coverage was at its lowest at the end of the dry growing season in 2002, indicating that drought limited plant growth. Vegetation cover recovered and consequently biomass increased during the following two growing seasons. There was two to ten times more plant biomass in 2004 than in 2003 (Table 3).
Even 20 to 30 years after ending all agricul- tural activities, the vegetation resembled more that of grass leys or meadows than mires. The plant biomass in 2004 was close to that of man- aged grass leys (Maljanen et al. 2001, 2004).
The predominant species, e.g. Deschampsia cespitosa, Juncus filiformis, Poa pratensis and Epilobium angustifolium, are known to favour
soils with high N-content (Jylhänkangas & Esala 2002). The secondary succession of abandoned fields to forested areas is known to be slow (Törmälä 1982, Jukola-Sulonen 1983, Kiirikki 1993). However, as described earlier, some scat- tered small birch trees and willow bushes were growing in the ditches.
Ecosystem and soil respiration rates The Reco and Rsoil rates during the growing season and winter were lower than those reported for cultivated peat fields in Finland (Maljanen et al. 2001, 2004) but higher than those from natu- ral peatlands (Alm et al. 1999a, Haapala et al.
2009). Measured Reco and Rsoil rates followed sea- sonal trends similar to those reported earlier for peatlands (e.g. Alm et al. 1997, Maljanen et al.
2001) and depended mainly on soil temperature and moisture. Both Reco and Rsoil rates were lower during the dry and warm growing season of 2002 than during the next, wetter growing season of 2003, indicating that low soil moisture limited ecosystem respiration in 2002, as has also been reported by Shurpali et al. (2009) for an organic soil covered with perennial grass. On the other hand, in 2004, the wet growing season and high WT levels also caused low respiration rates;
Rsoil was only half that in 2003. Mäkiranta et al.
(2009) showed an optimum WT for Rsoil in affor- ested agricultural peat soils and forestry-drained peatlands to be at about 60 cm. With higher or deeper water-table surface, wetness or drought apparently restricted soil respiration. High WT levels only slightly decreased Reco since the moist conditions favoured photosynthesis and plant growth, which was seen as higher GPP and greater plant biomass and coverage in 2004 than in the previous years. The Reco:Rsoil ratio was 2 in 2002, 2.4 in 2003 and 5.6 in 2004, showing that with increasing soil moisture Reco increased more than Rsoil. The autotrophic respiration can be estimated as Reco – Rsoil. The corresponding aver- age values of the autotrophic respiration rates were –504, –775 and –979 g CO2-C m–2 for the years 2002, 2003 and 2004, respectively. This indicates that good water availability favours autotrophic respiration more than heterotrophic respiration (Reco). The measured Rsoil in 2002
could even be overestimated since the cutting of above-ground plants was done in June 2002 and a lot of degradable root material was then left in the soil (e.g. Alm et al. 2007).
Gross primary production (GPP) and net ecosystem CO2 exchange (NEE)
The interannual variation in WT levels and asso- ciated variation in plant coverage and biomass explained the differences in the CO2 balance
b
0 200 400 600 800 1000 1200 1400 0
200 400 600 800 1000 1200 1400
AB1 AB2 AB3 AB4 AB5 0 500 1000 1500 2000 2500 3000
Measured RECO (mg CO2-C m–2 h–1)
Measured GPP (mg CO2-C m–2 h–1) Modelled GPP (mg CO2-C m–2 h–1)Modelled RECO (mg CO2-C m–2 h–1)
0 500 1000 1500 2000 2500 3000 a
Fig. 6. comparison of measured and modelled day- time (a) ecosystem respiration (Reco) and (b) gross pri- mary production (GPP) of abandoned agricultural soils (aB1–aB5). the line is indicating 1:1 agreement. Reco and GPP are all shown as positive values.
between the years (Fig. 5). GPP and Reco models (Eqs. 2 and 3) fit well with the data (Fig. 6).
During the first dry growing season, GPP at all sites was lower than during the following wetter growing seasons. The low WT level in 2002 resulted in low biomass production. The measured GPP values in the dry year 2002 were within the range measured in Finland for grass leys on peat soils with low management intensity (Maljanen et al. 2001, 2004), but in the follow- ing years they were 20%–40% higher than those for grass leys.
The annual net CO2 exchange (NEE) varied from slightly negative (i.e. uptake), at –8 g C m–2 yr–1 (wet year), to net emissions of 530 g C m–2 yr–1 (dry year). A similar climate-depend- ent trend has been found for perennial grass cultivated on boreal organic soil (Shurpali et al.
2009). In both cases, changes in GPP mostly depended on the hydrology of the soil. Zhou et al. (2011) further showed in a greenhouse experi- ment that the GPP of the perennial reed canary grass was sensitive to water availability. Alm et al. (1999b) also reported net CO2 losses from a natural peatland as a result of reduced photosyn- thesis during a dry summer. The results in this study and those of Shurpali et al. (2009) showed that respiration in drained organic soil can be lim- ited by low water availability. In wet years, the increase in GPP can more than compensate for the increase in C loss from increased soil respira- tion. However, if the soil water content is high enough, the respiration rate is reduced as a result of oxygen deficiency in the soil whereas GPP still increases, enhancing the net CO2 uptake.
The high GPP at sites AB1 and AB2 during the wet growing season could be associated with the Juncus species, in which aerenchyma allows for gas diffusion between shoots and roots (Din- smore et al. 2009), i.e. transport of oxygen to the roots when the WT level is high. According to Law et al. (2002), photosynthesis in boreal forest ecosystems during a cloudy day can be higher than on sunny days, which also supports our results of high GPP during the rainy growing season.
The average net CO2-C emissions during the first two years, excluding the net uptake during the wet growing season of 2004, were in the range reported for cultivated peatlands in
